Cretaceous faunas from Zululand and Natal, South Africa. The ammonite subgenus Hauericeras (Gardeniceras) Matsumoto & Obata, 1955

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1 Cretaceous faunas from Zululand and Natal, South Africa. The ammonite subgenus Hauericeras (Gardeniceras) Matsumoto & Obata, 1955 William James Kennedy 1 * & Herbert Christian Klinger 2 1 Department of Earth Sciences, Oxford University, South Parks Road, Oxford OX1 3AN, U.K. 2 Natural History Collections Department, Iziko South African Museum, P.O. Box 61, Cape Town, 8000 South Africa Received 6 September Accepted 14 December 2011 Four species of the distinctive desmoceratid ammonite Hauericeras (Gardeniceras) Matsumoto & Obata, 1955, are described from KwaZulu-Natal and the Eastern Cape Province: the type species, H. (G.) gardeni (Baily, 1855), from the Santonian to Lower Campanian, H.(G.) pseudoangustum Collignon, 1961, and H. (G.) madagascariense Collignon, 1961, from the Lower Campanian, and H. (G.) rembda (Forbes, 1846) from the upper Upper Maastrichtian. Keywords: ammonites, Hauericeras (Gardeniceras), Santonian, Campanian, Cretaceous, KwaZulu, Eastern Cape Province, South Africa. INTRODUCTION Hauericeras (Gardeniceras) gardeni (Baily, 1855) is one of the most distinctive and immediately recognizable ammonites from the Santonian Campanian Mzamba Formation of the coastal outcrops in the northeast of the Eastern Cape Province. In the cliff outcrops of the St Lucia Formation on the northwestern shores of False Bay, in KwaZulu-Natal, the species is abundant, sometimes occurring as imbricate stacks of individuals lying inclined at a low angle to bedding, and overlapping like roof tiles. The distinctive keel of the genus led Matsumoto (1938) to introduce a subfamily Hauericeratinae, supported in more extensive discussions by Matsumoto & Obata (1955) and Matsumoto et al. (1990); the subgenus Gardeniceras was introduced by Matsumoto & Obata (1955, p. 134) for the Fairly evolute and fairly widely umbilicate subgroup of Hauericeras and later (Matsumoto in Matsumoto et al. 1990, p. 456) redefined as a subgroup of Hauericeras in which ventral or ventrolateral riblets or nodes disappeared completely. Wright (1957, p. L371; 1996, p. 85) retained the subfamily, and defined it as follows (the phrase in italics was our addition to the 1996 account): Rather evolute to rather involute; whorl section high, with flat sides, venter rounded (at least initially), then typically fastigiate, and later with high sharp septicarinate keel. Smooth or with weak tubercles on the shoulders. Microconchs with lappets. Suture with suspensive lobe retracted or not. Origin is doubtful, either in Desmoceratinae close to point of origin of Muniericeratidae or Parapuzosia of Puzosiinae. Wright defined Hauericeras as having characters as for subfamily, and noted that Separation of Gardeniceras for the less involute forms seems unnecessary. Apart from Hauericeras, the genera In current geopolitical terminology Zululand and Pondoland now form parts of the provinces of KwaZulu-Natal and the Eastern Cape, respectively. For the sake of continuity we retain the names Zululand and Natal in the title of our series of systematic descriptions of the invertebrate faunas from these regions from 1975 onwards. *Authors for correspondence. jim.kennedy@oum.ox.ac.uk / hklinger@iziko.org.za Mossamedites Cooper, 2003a, and Oiophyllites Spath, 1953, have been referred to the Hauericeratinae. The latter is based on tiny desmoceratoidean nuclei and is best treated as a nomen dubium. In introducing his new genus Hauericeras, de Grossouvre (1894, p. 219) stated that Je donne ce nom à la série de formes se rattachant à Ammonites gardeni, but took Ammonites pseudogardeni Schlüter, 1872, p. 54, pl. 16, figs 3 6, as type species. Hauericeras (Hauericeras) pseudogardeni has been revised in detail by Kaplan & Kennedy (1995, p. 18, pls 1 4, pl. 5, figs 1 2; pl. 6, figs 1, 7; pls 7, 8) and Kaplan et al. (2006, p. 31, pl. 1, figs 4, 7, 9, 11 12). These authors redescribed the type material from Dülmen in Westphalia, and additional specimens from Braunschweig in Lower Saxony, Germany. From the material available it seems that internal moulds may be virtually smooth (but for constrictions), whereas the shell surface and composite moulds bear ribs/tubercles on the outer flanks and ventrolateral shoulders. But even some composite moulds are virtually smooth, although this might be due to postmortem effects. If this interpretation is accepted, variety nodatum of Schlüter (1899) and Hauericeras buszii Wegner, 1905 (p. 209, pl. 8, fig. 1a, b) are synonyms of pseudogardeni. The illustrations of the species in Müller & Wollemann (1906) of material from Braunschweig, included constricted ribbed/nodate phragmocones up to 160 mm diameter (1906, pl. 4, fig. 1; pl. 8, fig. 3) as well as smooth, delicately constricted juveniles. Matsumoto in Matsumoto et al (p. 451) thought Ammonites mengedensis Schlüter, 1876 (p. 154, pl. 40, fig. 9) might be the microconch of H. (H.) pseudogardeni, but this is a significantly older species (Kaplan & Kennedy 1994) and does not co-occur with H. (H.) pseudogardeni in any of the collections studied by Kaplan & Kennnedy, who were unable to conclusively demonstrate dimorphism. Hauericeras (H.) pseudogardeni is firmly dated as Upper Santonian to Lower Campanian in Western Europe. Matsumoto et al. (1990) and Kaplan & Kennedy (1995) suggested that Hauericeras (H.) antiquum Collignon, 1961 ISSN Palaeont. afr. (December 2011) 46:

2 (p. 75, text-fig. 12) known from a single specimen from the Lower Coniacian of Madagascar might be the stock ancestral to H. (H.) pseudogardeni, having the shell shape of Hauericeras plus delicate ventral ribs, but no constrictions. It is transitional to the Middle Turonian Puzosia (Puzosia) serratocarinata Kennedy & Cobban 1988 (p. 595, text-fig. 2; text-fig. 4, figs 1 3), from northern Mexico and Angola, the type species of Mossamedites Cooper, 2003 (p. 115) with a fastigiate venter but no well-differentiated keel. Gardeniceras first appears in the Middle Coniacian, with Ammonites lagarus Redtenbacher, 1973, from the Middle Coniacian of Austria and southeastern France (see revision in Kennedy in Kennedy et al. 1995, p. 397, pl. 4, fig. 17), and ranges to the upper Upper Maastrichtian, with Gardeniceras rembda (Forbes, 1846) (see revision in Kennedy & Henderson 1992, p. 408, pl. 6, figs 10 24; pl. 17, fig. 1; text-fig. 3H). Gardeniceras thus appears a little later than Hauericeras sensu stricto if antiquum is accepted as a Hauericeras: Middle versus Lower Coniacian. When compared to Hauericeras sensu stricto, Gardeniceras lacks outer flank and ventrolateral ribs/tubercles, has a keel that is present on the shell from an early ontogenetic stage, hollow when first secreted, but thereafter filled in by additional shell material so that it is solid over the phragmocone, as a result of which internal moulds of phragmocones have a rounded venter, rather than the sharply fastigiate venter with solid keel of H. (H.) pseudogardeni. The venter of internal moulds of Gardeniceras body chambers retains a rounded venter, with a keel that remains hollow. If Hauericeras and Gardeniceras are closely related, the stratigraphic evidence suggests the former gave rise to the latter during the Coniacian. Interestingly, the juvenile Mossamedites serratocarinatus figured by Kennedy (1988, pl. 2, figs 8 10) has the overall proportions of Gardeniceras. In conclusion, we accept Gardeniceras as a subgenus of Hauericeras in the absence of evidence to the contrary, but note that a keel has appeared more than once in the Desmoceratidae; in Damesites Matsumoto, 1942, Moremanoceras Cobbban, 1972, and in an as yet undescribed form from the Lower Cenomanian of Nigeria. It may well be that Hauericeras and Gardeniceras are homoeomorphs that acquired a keel independently. The only distinctive feature to justify Hauericeratinae is the presence of a keel, and as noted, this feature has appeared more than once in the evolution of the Desmoceratidae and we regard Hauericeratinae as unnecessary, and place Hauericeras in Puzosiinae contrary to the views of Matsumoto (1938, pp. 6 7) Crick (1907, p ; pl. 15, fig. 5) referred two poorly preserved fragments of compressed ammonites from the southern branch of the Manuan Creek in northern KwaZulu-Natal to Hauericeras. They are illustrated here as Fig. 10 I L, and appear to be fragments of an Albian or Cenomanian puzosiine, possibly Bhimaites Matsumoto, FIELD LOCALITIES Details of field localities are given by Kennedy & Klinger (1975); further descriptions of these localities are deposited in the Geological Collections, Oxford University Museum of Natural History, The Natural History Museum, London, and the Natural History Collections Department, Iziko South African Museum, Cape Town. SYSTEMATIC PALAEONTOLOGY Superfamily Desmoceratoidea Zittel 1895 Family Desmoceratidae Zittel, 1885 Subfamily Puzosiinae Spath, 1922 (= Hauericeratinae Matsumoto, 1938) Genus Hauericeras de Grossouvre, 1894 Schlueteria Rollier, 1922, p. 359, non Fritsch in Fritsch & Kafka, 1887, p. 33; Pseudogardenia Tomlin, 1930, p. 23. Type species Ammonites pseudogardeni Schlüter, 1872, p. 54, pl. 16, figs 3 6, by original designation by de Grossouvre, 1894, p Subgenus Gardeniceras Matsumoto & Obata, 1955 Type species Ammonites gardeni Baily, 1855, p. 450, pl. 11, fig. 3, by original designation by Matsumoto & Obata, 1955, p Hauericeras (Gardeniceras) gardeni Baily, 1855, Figs 1A C, 2, 3A D, 4A F, 5A H, 6, 7, 10M O 1855 Ammonites gardeni Baily, p. 450, pl. 11, fig. 3. non 1865 Ammonites gardeni Baily; Stoliczka, p. 61, pl. 33, fig. 4. (= H.(G.) angustum Yabe, non 1869 Ammonites Gardeni Baily; Favre, p. 12, pl. 4, fig. 1 (= Hauericeras (Gardeniceras) sulcatum (Kner, 1850)) Ammonites rembda Forbes; Griesbach, p. 63, pl. 3, figs 2 3.?non 1879 Ammonites Gardeni Baily; Whiteaves, p non 1890 Desmoceras gardeni (Baily); Yokoyama, p. 184, pl. 20, fig. 10 (=Hauericeras (Gardeniceras) angustum Yabe, 1904) Hauericeras gardeni (Baily); de Grossouvre, p. 219.?non 1895 Desmoceras Gardeni (Baily); Whiteaves, p non 1898 Hauericeras Gardeni (Baily); Kossmat, p. 123 (188), pl. 18 (24), fig. 7.?non 1903 Hauericeras gardeni (Baily); Whiteaves, p Hauericeras Gardeni (Baily); Woods, p. 332.?non1907 Hauericeras cf. Gardeni (Baily); Pervinquière, p. 166, pl. 7, fig. 1, 3 6.?1908 Desmoceras (Hauericeras) cf. Gardeni (Baily); Kilian & Reboul, p. 18. non 1913 Hauericeras gardeni (Baily); Nowak, p. 371, pl. 41, fig. 12; pl. 43, fig. 34; pl. 45, figs 44, 45 (= Hauericeras (Gardeniceras ) sulcatum (Kner, 1850) 1921 Hauericeras gardeni (Baily); Van Hoepen, p. 27, fig Hauericeras gardeni (Baily); Spath, p. 238, text-fig. A-1 to A Hauericeras gardeni, Baily sp.; Spath, p Hauericeras gardeni (Baily); Diener, p. 95 (pars) Hauericeras gardeni Baily; Besairie, p. 220, pl. 20, fig ISSN Palaeont. afr. (December 2011) 46: 43 58

3 A B C D E Figure 1. A C, Hauericeras (Gardeniceras) gardeni (Baily, 1855). The lectotype, BMNH C72219, the original of Baily 1855, p pl. 11, fig. 3a, b. D, E, Hauericeras (Gardeniceras) pseudoangustum Collignon, 1961, BMNH C35621, a paralectotype of Hauericeras (Gardeniceras) gardeni (Baily, 1855). Both specimens are from White-mens houses, coast of S. Africa near the Umzambani River, that is to say the Santonian to Lower Campanian Mzamba Formation at locality 1 of Kennedy & Klinger (1975), in Eastern Cape Province. Figures are Hauericeras Gardeni Baily; Basse, p. 23, pl. 4, figs 2 4, pl. 10, fig. 8; pl. 11, fig Hauericeras gardeni (Baily); Collignon, p. 17, pl. 3, fig Hauericeras Gardeni (Baily); Collignon, p. 74. non 1942 Hauericeras gardeni (Baily); Matsumoto, p. 25 (=Hauericeras (Gardeniceras) angustum Yabe, 1904).?non 1952 Hauericeras gardeni (Baily); Usher, p. 65, pl. 5, figs 1, 2; pl. 21, fig Hauericeras (Gardeniceras) gardeni (Baily); Matsumoto & Obata, p. 140 et seq.; text-figs Hauericeras gardeni (Baily); Collignon, p. 76, pl. 28; pl. 29; pl. 30, figs 1, 2; text-figs Hauericeras (Gardeniceras) gardeni Baily; Collignon, p. 66, pl. 539, fig Hauericeras cf. H. gardeni (Baily); Kennedy & Klinger in Kennedy, Kauffman & Klinger, p. 101, p. 6, fig Hauericeras gardeni (Baily); Kennedy & Klinger, p. 279, Hauericeras gardeni (Baily, 1855); Klinger & Kennedy, p. 80, text-figs 7A C, 8A. non 1979 Hauericeras (Gardeniceras ) gardeni (Baily); Summesberger, p. 133, pl. 6, fig. 27; text-fig Hauericeras gardeni (Baily); Immel, Klinger & Wiedmann, p. 16 (pars), pl. 6, figs 2, 3, non pl. 5, figs 1 4; non pl. 6, figs 1,4; non text-fig. 5. non 1982 Hauericeras (Gardeniceras) aff. gardeni (Baily); Renz, p. 106, pl. 35, figs 2 4. non 1987 Hauericeras gardeni (Baily); Immel, p Hauericeras gardeni (Baily, 1855); Matsumoto, Toshimitsu & Kawashita, p Hauericeras (Gardeniceras) cf. gardeni (Baily, 1855); Kennedy in Kennedy, Bilotte & Melchior, p b Hauericeras (Gardeniceras) gardeni (Baily1855); Cooper, p. 159, figs 6A D. ISSN Palaeont. afr. (December 2011) 46:

4 Figure 2. External suture of Hauericeras (Gardeniceras) gardeni (Baily, 1855), based on OUM KX Types Spath (1921, p. 238) refers to Baily s type (BM Geol. Soc. Coll., no ) and the larger of the two fragmentary cotypes (No ) from which Baily s figure of the suture line was taken, and subsequently (1922, p. 131) refers to BMNH as Baily s paratype. This does not constitute a valid lectotype designation in our view. Accordingly, we designate BMNH C72219 (Geological Society Collection no. R11370) lectotype. It is the original of Baily 1855, pl. 11, figs 3a,b, and is illustrated here as Fig. 1A C. The locality is given on an attached contemporary paper label as White-mens houses Umzambani River. Paralectotype BMNH C35621 (Geological Society Collection no. R11369) is figured here as Fig. 1D,E, and is referred to Hauericeras pseudoangustum Collignon, 1961, below. It has an associated grey-blue paper label: Ammonites gardeni nov. sp. Cretaceous Formation Amzamba River Port Natal. Presd. By Captain Garden D.B. p Paralectotype BMNH C35622 (Geological Society Collection no. R11370) has the same associated information, and is illustrated here as Fig. 3A,B; it provided the basis for Baily s illustration of the suture line of the species (1855, pl. 11, fig. 3c). Garden (1855, p. 453) gave the following account, About three miles to the southward of the river commence certain excavations in the cliffs, formed by the actions of the sea, and called by the natives Izinhuluzabalungu the houses of the white men so called probably from the caverns having once been occupied by shipwrecked sailors. This clearly identifies the locality of the types as locality 1 of Kennnedy & Klinger (1975), illustrated as text-figs 1 3 in Klinger & Kennedy (1980), the Izinhluzabalungu Caves of Griesbach (1871, fig. 5) and the Umzamba Cliff of Plows (1921), on the north side of the Mzamba estuary. Material BMNH C8516 and C18517 (both mentioned by Spath 1922, p. 130), C18518 (mentioned by Spath 1922, p. 130, figured by Matsumoto & Obata 1955, p. 141, text-fig. 10), C18519, C18520 (both mentioned by Spath 1922, p. 130, the latter figured by Matsumoto & Obata 1955, p. 140, text-fig. 9), C18521 and C18522 (both mentioned by Spath 1922, p. 130), C , C18528 (the original of Spath 1921, p. 238, fig. A1 7) (Fig. 3C), C18530, C18531 (the original of Spath 1921, p. 238, text-fig. A-8 (Fig. 3D herein), C18532, C18534, C18535 (mentioned by Spath 1922, p. 131), C18536, C18537 (mentioned by Spath 1922, p. 130), C18538, C18539, C18540 (mentioned by Spath 1922, p. 130), C18541, C , C All of these specimens are from the Mzamba Formation of locality 1 of Kennedy & Klinger (1975) although the details on associated labels vary slightly. OUM KX90, SAM-PCZ* all from locality 1 of Kennedy & Klinger (1975), where the species ranges from Santonian III to Campanian I (Klinger & Kennedy 1980, fig. 4). SAM BH9/120/22, BH9/124, BH9/127, 80, from the St Lucia Formation, Santonian III, in the BH9 borehole, 14 km west of Richards Bay in northern KwaZulu-Natal. OUM KX and SAM-PCZ* from the St Lucia Formation, Campanian I, at locality 14 of Kennedy & Klinger (1975) south of Mtubatuba, northern KwaZulu-Natal. OUM KX5353, 5356, 10838, 10839, 10841, 10843, 10845, SAM-PCZ* from the St Lucia Formation, Santonian I, locality 17 of Kennedy & Klinger (1975), southeast of Mtubatuba, northern KwaZulu-Natal. OUM KX4970a b, SAM-PCZ* from the St Lucia Formation, Campanian I at locality 105 of Kennedy & Klinger (1975; see also Klinger & Kennedy 1980a, fig. 130) on the southeastern shores of False Bay, lake St Lucia, northern KwaZulu- Natal. OUM KX12818, , 12880, 12881, , 13037, , 13078, SAM-PCZ* from locality 74 of Kennedy & Klinger (1975), Die Rooiwalle, northwestern False Bay, Lake St Lucia, northern KwaZulu-Natal. The species has been collected in situ from beds 2 22 of the section, spanning the Santonian I Campanian I interval. *Unregistered. 46 ISSN Palaeont. afr. (December 2011) 46: 43 58

5 C A D B Figure 3. A D. Hauericeras (Gardeniceras) gardeni (Baily, 1855). A, B, paralectotype BMNH C35622, the original of Baily, 1855, pl. 11, fig. 3c (suture), from White-mens houses, coast of S. Africa near the Umzambani River, that is to say the Santonian to Lower Campanian Mzamba Formation at locality 1 of Kennedy & Klinger (1975), in Eastern Cape Province. C, BMNH C18528, the original of Spath 1921, p. 238, text-fig. A-1-7, 9. D, BMNH C18531, the original of Spath 1921, p. 238, text-fig. A-8, from the Upper Santonian to Lower Campanian Mzamba Formation at locality 1 of Kennedy & Klinger (1975), in Eastern Cape Province. Figures are 1. Dimensions D Wb Wh Wb:Wh U KX (100) 15.2(20.3) 25.5 (34.0) (41.7) C (100) 16.4 (21.0) 29.0 (37.2) (37.9) C (100) 16.2 (19.9) 30.4 (37.4) (35.3) KX (100) 16.1 (19.6) 29.2 (35.5) (40.1) C (100) 18.0 (20.7) 32.2 (37.0) (40.6) KX (100) 16.8 (18.8) 31.7 (35.5) (41.5) C ( (19.8) 33.0 (35.4) (37.2) C (100) 19.0 (20.3) 31.6 (33.8) (41.3) C (100) 21.4 (20.2) 37.4 (35.3) (38.7) KX (100) 20.8 (18.7) 39.5 (35.5) (42.6) KX (100) 20.1 (16.1) 42.5 (34.1) (42.6) C (100) 22.8 (17.5) 42.6 (32.6) (44.5) C (100) 28.0 (21.4) 43.3 (33.1) (41.0) C (100) 22.2 (16.8) 42.3 (32.1) (43.1) Description The inner septate whorls are well represented by the lectotype, BMNH C72219 (Baily 1855, p. 456, pl. 11, fig. 3a, b; Fig. 1A C herein) which is partially exfoliated, retaining original aragonitic shell layers. There are indications of the former presence of at least one further whorl. Coiling is very evolute, the umbilicus very broad and shallow, comprising 37.9% of the diameter, with 41% of the previous whorl covered. The low umbilical wall is flattened and outward-inclined with a shallow groove just below the umbilical shoulder. The umbilical shoulder is sharp on the penultimate whorl, becoming rounded on the outer whorl. The whorl section is compressed, with a whorl breadth to height ratio of 0.57, the greatest breadth low on the flanks, the inner flanks flattened and feebly convergent, the outer flanks convex and converging to the acutely fastigiate venter. A strong siphonal keel is present where shell is preserved, but absent on the internal mould. There is no ornament preserved on the surface of the exfoliated shell. Part of a single constriction is visible on the internal mould on one flank. The delicate growth lines shown in Baily s figure are not detectable. Paralectotype BMNH C35622 is the basis of Baily s figure of the suture (1855, pl. 11, fig. 3c; Matsumoto & Obata ISSN Palaeont. afr. (December 2011) 46:

6 A B C D E F Figure 4. Hauericeras (Gardeniceras) gardeni (Baily, 1855). A C, OUM KX10843; D F, OUM KX10840, from the St Lucia Formation, Santonian III, locality 17 of Kennedy & Klinger (1975), southeast of Mtubatuba, northern KwaZulu-Natal. Figures are , text-fig. 8; Fig. 3A,B). It comprises fragmentary inner whorls and a 180 sector of outer whorl; the estimated maximum diameter is 130 mm. The adapertural 90 sector is body chamber. The specimen retains partially exfoliated shell in places. Constrictions are not detectable on the early phragmocone whorls. A weak constriction is present at the adapertural end of the phragmocone, and there is a single constriction on the body chamber fragment. The constrictions are concave, projecting strongly forwards on the ventrolateral shoulders and venter. The constrictions are far less conspicuous than is suggested by the sketch in Matsumoto & Obata (1955, text-fig. 8a). BMNH C18518 (Fig. 7) is the thickest form mentioned by Spath (1922, p. 130), and figured by Matsumoto & 48 ISSN Palaeont. afr. (December 2011) 46: 43 58

7 A B C D E F G H Figure 5. Hauericeras (Gardeniceras) gardeni (Baily, 1855). A, B, OUM KX10844; C, D, OUM KX5350; E H, OUM KX5349, all from the St Lucia Formation, Santonian III, locality 17 of Kennedy & Klinger (1975), southeast of Mtubatuba, northern KwaZulu-Natal. Figures are 1. Obata (1955, text-fig. 10). The specimen has a maximum preserved diameter of mm, and retains extensive areas of shell. Coiling is very evolute. The wide, shallow umbilicus comprises 41% of the diameter, the umbilical wall low, the umbilical shoulder very narrowly rounded. The whorl breadth to height ratio is 0.65, the flanks feebly convex, and feebly converging, the greatest breadth just outside the umbilical shoulder. The ventrolateral shoulders are broadly rounded, the venter acutely fastigiate with a strong sharp keel where shell is preserved on both phragmocone and body chamber, and on the internal mould of the body chamber. The internal mould of the phragmocone lacks a keel. The shell surface is smooth on the phragmocone, apart from delicate growth lines that ISSN Palaeont. afr. (December 2011) 46:

8 Figure 6. Hauericeras (Gardeniceras) gardeni (Baily, 1855). OUM KX 90, from the Mzamba Formation, Santonian III, at locality 1 of Kennedy & Klinger (1975), in Eastern Cape Province. Figures are 1. are feebly prorsiradiate on the inner flank, concave on the middle to outer flank and projected strongly forwards on the ventrolateral shoulders and venter, indicating the presence of a long ventral rostrum at the aperture. The adapertural 240 sector is body chamber, with four constrictions that become increasingly prominent as size increases. They are concave across the flanks and project strongly forwards on the ventrolateral shoulders and venter. The adapertural edge of the constrictions is marked by a feeble collar-rib. The final constriction appears to be just before a damaged apertural margin that bears a ventral rostrum. OUM KX 90 (Fig. 6) is 120 mm in diameter, and retains extensive traces of partially exfoliated shell. Six constrictions are detectable on both the outer whorl and the penultimate whorl. Very well-preserved body chambers from locality 17 (Figs 4A F, 5A H) show the presence of strong constrictions, three per half whorl, with an adapical collar-rib on the internal mould. There is little or no indication of the constrictions where shell is preserved on the opposite flank of the same individual (OUM KX10843: compare Figs 4A and C; OUM KX5349: compare Figs 5F and 5G). An unexplained feature of partially exfoliated specimens is particularly well-shown by BMNH C18520 (Fig. 10M O, the original of Spath 1922, p. 130, and Matsumoto & Obata 1955, text-fig. 9). The translucent layers of replaced shell material show a pattern of deeply concave, narrow dark bands (much more deeply concave than the constrictions) separated by wider white bands. This feature is largely within the shell material, being scarcely indicated on the surface, as can be seen from the whitened and unwhitened picture of the specimen (Fig. 10N, uncoated; Fig. 10O, coated with ammonium chloride). These patterns presumably relate to the accretion pattern of the shell material; similar structures are shown by BMNH C18535 and C A keel is present on phragmocones and body chambers with shell preserved, and on internal moulds of body chambers. The keel was thus initially hollow and open to the interior of the shell, and thereafter infilled with shell material and solid on the phragmocone, but remained hollow on the body chamber. (See also Matsumoto & Obata 1955, text-figs 8 10; Klinger & Kennedy 1977, figs 7a,c, 8a.) The suture (Fig. 2) is deeply incised, with asymmetrically bifid E/A and A/U2, deep asymmetrically trifid A (=L) and U2, the suspensive lobe strongly retracted. Discussion Hauericeras (Gardeniceras) gardeni, as interpreted from the lectotype, paralectotype BMNH C35622 and the abundant Mzamba material referred to the species, has a compressed whorl section with whorl breadth to height ratios 50 ISSN Palaeont. afr. (December 2011) 46: 43 58

9 A B C Figure 7. Hauericeras (Gardeniceras) gardeni (Baily, 1855). BMNH C18518, the original of Spath 1922, p. 130 and Matsumoto & Obata, 1955, text-fig. 10, from the Upper Santonian to Lower Campanian Mzamba Formation at locality 1 of Kennedy & Klinger (1975), in Eastern Cape Province. Figures are 1. of , the majority falling between 0.53 and 0.57, the umbilicus comprising between 35.3% and 47.0% of the diameter, the majority falling between 37.2% and 44.5%. Concave growth lines are only occasionally discernible, perhaps as a result of the partial exfoliation of most specimens. Concave constrictions are weak or imperceptible on the surface of the shell, but conspicuous on the internal mould, and are markedly strengthened on the body chamber, with an associated adapertural collar rib. On this basis, H. (G.) gardeni can be distinguished from the closely allied H. (G.) angustum Yabe, 1904 (p. 33, pl. 5, figs 5, 6) which was carefully revised by Matsumoto & Obata (1955, p. 137, pl. 24, fig. 6; pl. 28, figs 1, 2; pl. 29, figs 1 5; text-figs 5, 7) and Matsumoto et al. (1990, p. 443, textfigs 2 7). On the basis of the observations of these authors, and specimens in the Natural History Museum, London (BMNH C , mentioned by Spath, 1922 p. 131) H. (G.) angustum has sinuous growth lines and constrictions that are concave on the innermost flank, convex on the inner to middle flank, and concave on the outer flank. Body chambers of gardeni have strengthened ribs on the body chamber, with a collar rib; they are weak on body chambers of angustum, some specimens of which develop what appear to be coarse inner flank ribs on the body chamber (Matsumoto & Obata 1955, pl. 28, fig. 2). Dimorphism has been recognized in H. (G.) angustum. Matsumoto et al. (1990, text-fig. 2) figured what they regarded as a macroconch 225 mm in diameter with a sinuous, biconcave mouth border (1990, text-fig. 3) and microconchs, the best preserved of which (1990, textfigs 6, 7) is 113 mm in diameter, with a long lappet at mid-flank, and longer ventral rostrum. Matsumoto et al. (1990, p. 451) thought that BMNH C18518, figured here as Fig. 7, was the macroconch of H. (G.) gardeni. This specimen is 135 mm in maximum preserved diameter, with an estimated 230 sector of body chamber, what appears to be a long ventral rostrum, but no trace of a lappet. Matsumoto et al. (1990, p. 452) then proposed that the two large figured specimens of H. (G.) gardeni from Madagascar figured by Collignon (1961, pls 28, 29) were macroconchs. These appear to lack the final sector of body chamber and are preserved to diameters of 183 and 195 mm. They are far larger than any material from Mzamba or KwaZulu-Natal that we have seen, and we are unable to confirm dimorphism in the material before us. Hauericeras (Gardeniceras) pseudoangustum Collignon, 1961 (p. 83, text-fig. 18) is smooth, with weak sinuous constrictions that are restricted to the body chamber. Hauericeras (Gardeniceras) madagascariense Collignon, ISSN Palaeont. afr. (December 2011) 46:

10 Figure 8. Hauericeras (Gardeniceras) madagascariense Collignon, SAM-7043, from the uppermost beds of the Lower Campanian of the Mzamba Formation at locality 1 of Kennedy & Klinger (1975), in the Eastern Cape Province. Figure is (p. 81, pl. 31, fig. 1; pl. 32, fig. 1; text-figs 15 17) differs from H. (G.) gardeni in its much broader whorl section, with whorl breadth to height ratios of up to 0.76, and sinuous, biconcave constrictions. If the strength and course of the constrictions is regarded as a key feature, then material with sinuous rather than simple concave constrictions must be excluded from H. (G.) gardeni. These include material from Tunisia (Pervinquière 1907), Austria (Summesberger 1979; Immel et al. 1982; Immel 1987) and Venezuela (Renz 1982). Occurrence In South Africa, H. (G.) gardeni first appears in Santonian I and ranges into Campanian I of Kennedy & Klinger (1975), with records from the Mzamba Formation of the coastal outcrops in northeastern Eastern Cape Province, and the St Lucia Formation of northern KwaZulu-Natal. In Madagascar it ranges from Upper Santonian through all of the Lower Campanian. There are also records from the Santonian of southeastern France and Angola. Records of H.(G.) gardeni from the Santonian Campanian of south India and Vancouver Island, British Columbia, Canada are best referred to H.(G.) angustum as pointed out by Matsumoto & Obata (1955, p. 144) and Matsumoto (1959, p. 25) Hauericeras pseudoangustum Collignon, 1961, Figs 1D,E; Hauericeras (Gardeniceras) pseudoangustum Collignon, p. 83, text-fig. 18 Type The holotype, by original designation, is no in the collections of the Laboratoire de Paléontologie of the Muséum National d Histoire Naturelle, Paris, the original of Collignon text-fig. 18, from the Lower Campanian Zone à Anapachydiscus arrialoorensis (= the Zone à Menabites boulei et Anapachydiscus arrialoorensis of 52 ISSN Palaeont. afr. (December 2011) 46: 43 58

11 A B Figure 9. Hauericeras (Gardeniceras) pseudoangustum Collignon, The holotype, no in the collections of the Laboratoire de Paléontologie of the Musém d Histoire Naturelle, Paris, the original of Collignon. 1961, text-fig. 18, from the Lower Campanian Zone à Anapachydiscus arrialoorensis (= the Zone à Menabites boulei et Anapachydiscus arrialoorensis of Collignon, 1969) of Iampolypoly-Antsirasira-Behamotra (Belo sur Triribihina), Madagascar. Figures are 1. Collignon 1969) of Iampolypoly-Antsirasira-Behamotra (Belo sur Tsiribihina), Madagascar, illustrated here as Fig. 9. Material BMNH C35621 (Fig. 1D,E), a paralectotype of Ammonites gardeni Baily, 1855, from the Mzamba Formation at the mouth of the Mzamba River, Eastern Cape Province, corresponding to locality 1 of Kennedy & Klinger (1975). Description BMNH C35621 is an internal mould of a body chamber fragment with a maximum preserved whorl height of 63.2 mm and a whorl breadth to height ratio of There are traces of a single feeble constriction that is convex on the inner flank, concave on the outer flank, and projected strongly forwards on the outermost flank and ventrolateral shoulder. The estimated total original diameter of the present specimen is 170 mm. Discussion Whorl section and course of the single constriction separate this specimen from Hauericeras (Gardeniceras) gardeni, but correspond to that of Hauericeras (Gardeniceras) pseudoangustum, to which it is referred. Collignon s (1961, pp ) description leaves no doubt as to the affinities of the specimen: Coquille très plate, discoidale, à tours élevés, se recouvrant d un peu plus de 2/5; flancs plats, parallèles au voisinage de l ombilic et jusque vers leur milieu, puis s abaissent doucement vers la région externe arrondie d abord, puis devenant ogivale avec carène. Versant ombilical relativement élevé, vertical, limité par une arrête mousse. Ombilic moyennement large dégeant les tours internes plats, en escalier. Aucune ornamentation; seule un constriction est visible très nettement au début de la chambre d habitation, immédiatement en avant de la dernière cloison. Elle est légèrement tordue en S étiré, très allongé. Sur le plus grand exemplaire il y a 3 constrictions visibles, toutes sur la chambre d habitation. Occurrence Uppermost Santonian or Lower Campanian, Mzamba Formation at locality 1 of Kennedy & Klinger (1975); Lower Campanian Zone à Menabites boulei et Anapachydiscus arrialoorensis of Madagascar. ISSN Palaeont. afr. (December 2011) 46:

12 B C A H D E F G I J K L M N O Figure 10. A H. Hauericeras (Gardeniceras) rembda (Forbes, 1846). A, BMNH C51023, a paralectotype; G, H, BMNH C51024, the lectotype. Both specimens are from the Upper Maastrichtian Valudavur Formation of Pondicherry, south India. B E, OUM KX6973, from the St Lucia Formation, Maastrichtian II, upper Upper Maastrichtian Inoceramus ianjonaensis Zone of locality 20 of Kennedy & Klinger (1975), east-southeast of Mtubatuba, northern KwaZulu-Natal. F, OUM KX8906, from the St Lucia Formation, Maastrichtian III, upper Upper Maastrichtian of locality 128 of Kennedy & Klinger (1975), Lake St Lucia, northern KwaZulu-Natal. I L, indeterminate Puzosiinae. I, J, BMNH C18275, the original of Hauericeras sp. of Crick, 1907, p. 242, pl. 15, fig. 5; K, L, BMNH C18276, the original of Hauericeras sp. of Crick, 1907, p Both specimens were described as being from the southern branch of the Manuan Creek. M O, Hauericeras (Gardeniceras) gardeni (Baily, 1855). BMNH C18520, the original of Spath 1922, p. 130 (table) and Matsumoto & Obata, 1955, text-fig. 9, from the Upper Santonian to Lower Campanian Mzamba Formation at locality 1 of Kennedy & Klinger (1975), in Eastern Cape Province. The specimen is uncoated in Figs M and N; the latter shows the distinctive colour banding within the partially exfoliated shell material. Figures A, D, E are 2; B, C, F O are ISSN Palaeont. afr. (December 2011) 46: 43 58

13 Figure 11. External suture of Hauericeras (Gardeniceras) rembda (Forbes, 1846), based on OUM KX6937. Hauericeras (Gardeniceras) madagascariense Collignon, 1961, Fig Hauericeras (Gardeniceras) madagascariense Collignon, p. 81, pl. 31, fig. 1; pl. 32, fig. 1; text-figs Hauericeras (Gardeniceras) madagascariense Coll.; Collignon, p. 66, pl. 539, fig b Hauericeras madagascariense Collignon, 1961; Klinger & Kennedy, p. 219, text-fig. 5B.?1996 Hauericeras (Gardeniceras) gardeni (Baily, 1855); Cooper & Greyling, p. 23, fig. 7m n. Type The holotype by original designation, is the orignal of Collignon, 1961, pl. 31, fig. 1, no in the collections of the Muséum National d Histoire Naturelle, Paris, from the Lower Campanian Zone à Anapachydiscus arrialoorensis (= the Zone à Menabites boulei et Anapachydiscus arrialoorensis of Collignon, 1969) of Iampolypoly-Antsirasira- Behamotra (Belo sur Tsiribihina), Madagascar. Material SAM-7043 from the Mzamba Formation, probably bed A15 of Klinger & Kennedy (1980, text-fig. 4) (Bed T2 of Gevers 1923; see also Gevers 1977) at locality 1 of Kennedy & Klinger (1875), Eastern Cape Province. OUM KX1781, from the St Lucia Formation, Campanian II at locality 109 of Klinger & Kennedy (1975) on the southwestern corner of the Nibela Peninsula, Lake St Lucia, northern KwaZulu-Natal. Description SAM-7043 (Fig. 8) is a large specimen (D = c.160 mm) embedded in a concretion with specimens of Baculites sulcatus Baily, 1855 on the reverse side (Klinger & Kennedy 1997, fig. 64a). Part of the last septum is exposed. The body chamber occupies a sector of slightly more than 180. Four concave constrictions are visible on the body chamber; the most prominent one near the adapical part. The umbilicus is wide, c. 45 % of the total diameter) OUM KX1781 is an internal mould of a phragmocone 190 mm in diameter, the umbilicus shallow, comprising 47.5% of the diameter, with a low, feebly convex, outward-inclined umbilical wall and narrowly rounded umbilical shoulder. The whorl breadth to height ratio is 0.64, the inner and middle flanks flattened and feebly convergent, the outer flanks broadly rounded, converging to the narrowly rounded, arched venter of the internal mould. This is smooth, with no indication of constrictions. In places, traces of recrystallized shell on the venter demonstrate the presence of a sharp siphonal keel. The suture is very deeply and intricately incised, the saddles narrow-stemmed, bifid, with deep asymmetrically trifid A (=L). Discussion Hauericeras (Gardeniceras) madagascariense differs from Hauericeras (Gardeniceras) gardeni in its greater size, broader whorls, slightly larger umbilical diameter and feebly convex rather than narrowly rounded venter. The constrictions, well seen in SAM-7043, are concave, rather than sinuous as they are in Hauericeras pseudoangustum. Occurrence. According to Collignon (1961, p. 82) H. (G.) madagascariense is restricted to the upper part of the Lower Campanian, zone of Anapachydiscus arrialoorensis redefined as the upper Lower Campanian Zone à Menabites boulei et Anapachydiscus arrialoorensis of Collignon, 1969 of Madagascar. Campanian II of KwaZulu-Natal. At Mzamba in the Eastern Cape, it has only been found in the topmost beds of the Mzamba Formation, above the level of abundant H. (G.) gardeni. Hauericeras rembda (Forbes, 1846), Figs 10A H, Ammonites Rembda Forbes, p. 111, pl. 7, fig Ammonites Durga Forbes, p. 104, pl. 7, fig. 11. non 1871 Ammonites rembda Forbes; Griesbach, p. 63, pl. 3, figs 2 3 (= Hauericeras gardeni)?1978 Hauericeras sp. cf. H. (Gardeniceras) rembda (Forbes); Matsumoto, Okada & Sakurai, pp , fig Hauericeras rembda (Forbes, 1846); Kennedy & Henderson, p. 408, pl. 6, figs 10 24; pl. 17, fig. 1; text-fig. 3H (with full synonymy). Types The lectotype, by the subsequent designation of Matsumoto & Obata (1955, p. 145), is BMNH C51024, the ISSN Palaeont. afr. (December 2011) 46:

14 original of Forbes 1846, pl. 7, fig. 3; paralectotypes are BMNH C51023 and C The figured syntype of Ammonites Durga Forbes, 1846, p. 104, pl. 7, fig. 11 is BMNH C All of these specimens are from the Upper Maastrichtian Valudavur Formation of Pondicherry, south India. Material OUM KX6973, from the St Lucia Formation, Maastrichtian II, upper Upper Maastrichtian Inoceramus ianjonaensis Zone of locality 20 of Kennedy & Klinger (1975), ESE of Mtubatuba, northern KwaZulu-Natal. OUM KX8906, from the St Lucia Formation, Maastrichtian III, upper Upper Maastrichtian of locality 128 of Kennedy & Klinger (1975), Lake St Lucia, northern KwaZulu-Natal. Description OUM KX6937 (Fig. 10B E) is a well-preserved phosphatized phragmocone fragment of a 60 whorl sector with a maximum preserved whorl height of 15.3 mm and a whorl breadth to height ratio of 0.64, the greatest breadth well below mid-flank. The inner and middle flanks are feebly convex, the outer flank and ventrolateral shoulders converge to the broadly rounded venter. The surface of the internal mould is smooth. A sector of a solid calcite replaced siphonal keel is present. The suture is deeply incised, with a large trifid A (= L) (Fig. 11). OUM KX8906 (Fig. 10F) is a composite mould of a 120 sector of two successive whorls with a maximum preserved whorl height of 14.8 mm. Coiling is very evolute, the umbilicus broad and shallow, with a feebly convex outward-inclined umbilical wall and more narrowly rounded umbilical shoulder. The specimen is partially embedded in matrix, but the whorl section and proportions correspond to those of the previous specimen. A poorly preserved constriction is present toward the apertural end of the fragment. It is markedly prorsiradiate, feebly sinuous (?) and strongly projected forwards on the ventrolateral shoulders and venter. Discussion The type material of Hauericeras rembda was revised by Kennedy & Henderson (1992, p. 408, pl. 6, figs 10 24; pl. 17, fig. 1; text-fig. 3H). Given the Upper Maastrichtian horizon of the present material, the whorl section, and apparent course of the constriction on OUM KX 8906, reference to rembda seems appropriate; the lectotype and largest paralectotype are illustrated here as Fig. 10A, G, H for comparison. The other Maastrichtian species, Hauericeras sulcatum (Kner, 1848) (see revision in Kennedy & Summesberger 1987, p. 27, pl. 1, figs 1 7; pl. 13, fig. 2) has six or seven constrictions per whorl, and these are concave. The two specimens from the type locality of the Mzamba Formation described by Griesbach (1871, p. 63, pl. 3, figs 2 3) as Ammonites rembda do not belong here and are probably H. (G.) gardeni. We have not been able to examine Griesbach s material it is presumed to have been destroyed during aerial bombardment in World War II. (Klinger 1995). Occurrence Upper Maastrichtian of India and northern KwaZulu- Natal. Maastrichtian of Madagascar. Kennedy acknowledges the support of the staff of the Geological Collections, Oxford University Museum of Natural History, and the Department of Earth Sciences, Oxford, and the financial assistance of the Oppenheimer Fund (Oxford). Klinger acknowledges financial support from the NRF (South Africa) and assistance from staff of the Natural History Collections Department, Iziko South African Museum, Cape Town. For access and logistic support in the St Lucia-False Bay Lakes region of KwaZulu-Natal we thank the staff of the isimangaliso Wetland Park Authority (Formerly Greater St Lucia Wetland Park Authority). ABBREVIATIONS Institutional BMNH The Natural History Museum, London. OUM Oxford University Museum of Natural History. SAM Natural History Collections Department, Iziko South African Museum, Cape Town. Anatomical [Dimensions are given in millimetres] D diameter Wb whorl breadth Wh whorl height U umbilicus c costal dimension ic intercostal dimension (Figures in brackets are dimensions as a percentage of the diameter.) The suture terminology is that of Korn et al. (2003): E external lobe A adventive lobe (= lateral lobe, L, of Kullmann & Wiedmann 1970) U umbilical lobe I internal lobe. REFERENCES BAILY, W.H Description of some Cretaceous fossils from South Africa. Quarterly Journal of the Geological Society of London 11, BASSE, E Monographie paléontologique du Crétacé de la Province de Maintirano. Mémoire géologique du Service des Mines, Madagascar 1931, BESAIRIE, H Recherches géologiques à Madagascar. Contribution à l étude des ressources minérales. Bulletin de la Société d Histoire naturelle de Toulouse 60, COBBAN, W.A New and little-known ammonites from the Upper Cretaceous (Cenomanian and Turonian) of the Western Interior of the United States. 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